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Class Aves (birds) II Respiratory system Circulatory system Digestion and excretion Metabolism Thermoregulation Senses Intelligence Communication Migration Dispersal Reproduction How to identify birds Lung tissue is denser than mammalian lung tissue Conchae warm and clean air on intake and retain moisture on exhalation Air sacs occupy up to 20% of the body cavity Class Aves (birds) II One-way flow of (always oxygenated) air over bird lungs more efficient than two-way flow in mammalian lungs 4-chambered heart (like mammals and crocodiles) Respiratory system Circulatory system Digestion and excretion Metabolism Thermoregulation Senses Intelligence Communication Migration Dispersal Reproduction How to identify birds Circulation • 4-chambered heart • largest hearts relative to body size – can be twice the size of mammalian hearts • heart size correlated with speed of flying, diving, altitude, and cold climates • can more fully drain ventricle compared to mammals • high blood pressure – subject to heart failure in times of stress 1 Class Aves (birds) II Respiratory system Circulatory system Digestion and excretion Metabolism Thermoregulation Senses Intelligence Communication Migration Dispersal Reproduction How to identify birds mouth crop – storage chamber - horny tongue - few salivary glands short pharynx esophagus – long and elastic stomach - proventriculus (enzymes) - gizzard (grinds food) Fig. 39.11 Page 891 Fig. 39.11 Page 891 Owls, hawks and kingfishers regurgitate bones, fur and feathers Gizzard muscular chamber lined with rough keratin contains sandy grit (ingested by the bird) best developed in seed-eating birds pancreas small intestine pellets are formed in the proventriculus heart liver gizzard 2 duodenum small intestine caeca – fermentation chambers large intestine cloaca receives ureters, genital ducts & large intestine reabsorbs water Fig. 39.11 Page 891 Fig. 39.11 Page 891 Excretory System Nitrogenous waste excreted as uric acid • excrete uric acid into cloaca • excess water reabsorbed in cloaca • forms white paste and mixes with feces Excess salt E lt secreted t d by b special i l salt lt glands l d • special salt glands above each eye secrete a high concentration of salt • salt solution excreted via internal or external nostrils Class Aves (birds) II Respiratory system Circulatory system Digestion and excretion Metabolism Thermoregulation Senses Intelligence Communication Migration Dispersal Reproduction How to identify birds Metabolism • High resting metabolic rate (songbirds have the highest BMRs of all vertebrates) • During flight, metabolic rate 10X BMR • However, However flight is energetically very efficient – a song bird flying 1 km expends <1% of the energy a mouse would use to run the same distance. 3 Birds have high basal metabolic rates Humans BMR for a 70 kg male is about 1560 – 1680 kcal/day 23 kcal/kg/day Mass-specific basal rate of metabolism as a function of body mass Species Mass (gms) kcal/kg/day Trumpeter Swan 8900 47 Brown Pelican 3500 75 Common Raven 850 108 American Kestrel 110 157 White-crowned Sparrow 27 324 House Wren 11 589 Rufous Hummingbird 3.5 1600 O2 consumption as a function of body weight all birds nonpasserines flying fox flying fox fruit pigeon mammals Class Aves (birds) II Respiratory system Circulatory system Digestion and excretion Metabolism Thermoregulation Senses Intelligence Communication Migration Dispersal Reproduction How to identify birds 4 Endothermic vs ectothermic Endothermic vs ectothermic • Endothermic - use their own body heat to maintain a warm, constant body temperature (birds and mammals) • Endothermic - use their own body heat to maintain a warm, constant body temperature (birds and mammals) • Ectothermic – use environmental temperature to regulate body temperature (invertebrates, fish, amphibians, reptiles) Thermoregulation 1. Heat production – The primary means for increasing heat production for birds is shivering. Thermoregulation 2. Evaporation – birds lack sweat glands so evaporation can occur only via respiratory system: - panting - gular flutter The large flight muscles (pectoralis) as well as the leg muscles play an important role in generating heat by shivering. Thermoregulation 3. Feathers - number of feathers varies seasonally; more during the winter than the summer Thermoregulation 4. Posture to reduce heat loss: - withdraw feet into plumage - tuck head & neck under wing - position of feathers controlled by dermal muscles 5 Thermoregulation 4. Posture to reduce heat loss: - withdraw feet into plumage - tuck head & neck under wing Thermoregulation 5. Legs & Feet • limited heat loss in cold weather because of thermal counter-current to increase heat loss: - extend wings Counter current heat exchange Thermoregulation 5. Legs & Feet • limited heat loss in cold weather because of thermal counter-current • can radiate heat during hot weather: increased blood flow Thermoregulation 5. Legs & Feet • limited heat loss in cold weather because of thermal counter-current • can radiate heat during hot weather: increased blood flow • storks & vultures defecate on legs to increase heat loss by evaporation Thermoregulation 6. Social Behavior - Communal roosts, e.g., Willow Ptarmigan can burrow in the snow up to 21 hrs a day in cold weather. 6 Class Aves (birds) II Summary of Thermoregulation 1. Heat production – shivering 2. Evaporation - gular flutter & panting 3. Feathers - number of feathers varies seasonally - position of feathers controlled by dermal muscles 4. Posture - to reduce heat loss: withdraw feet; tuck head under wing - to increase heat loss: extend wings 5. Legs & Feet - thermal counter-current - increased blood flow in hot weather 6. Social Behavior - communal roosts - huddling Birds have excellent vision! Birds have good hearing abilities. Respiratory system Circulatory system Digestion and excretion Metabolism Thermoregulation Senses Intelligence Communication Migration Dispersal Reproduction How to identify birds Vision • • • • Vision is the dominant sense in birds Eyes are large - must turn their heads Birds can see 2 – 3 x better than humans Color vision well-developed in diurnal birds - many cones • Nocturnal birds have few cones and many rods • Most birds can see UV • Some birds can see polarized light Hearing ranges of birds: For most birds, hearing is best at frequencies of 1 - 4 kHz (humans: 0.5 – 5 kHz), but some birds are able to detect higher frequencies (up to 10 - 12 kHz). The ability of birds to discriminate differences in the frequency of sounds & to detect gaps between sounds is generally similar to humans. 7 Owls note the differences in intensity and timing of sounds between the two ears. Some owls have asymmetrical ear openings, allowing them to accurately note differences in the intensity and timing of sounds in both the horizontal (azimuth) and vertical (elevation) planes. Some owls have asymmetrical ear openings, allowing them to accurately note differences in the intensity and timing of sounds in both the horizontal (azimuth) and vertical (elevation) planes. Barn owls can capture a mouse in total darkness based on sound alone. Woodpeckers can hear grubs moving below the bark of trees and can detect low-frequency sounds that precede earthquakes. th k Birds lack conspicuous external ears. Instead, they have specialized feathers called auriculars or ear coverts. 8 Olfaction Except for flightless birds, ducks and vultures, birds have a poor sense of smell. “Dumb as a dodo” Dodo (Raphus cucullatus) Class Aves (birds) II Respiratory system Circulatory system Digestion and excretion Metabolism Thermoregulation Senses Intelligence Communication Migration Dispersal Reproduction How to identify birds Mauritius The great auk (Pinguinis impennis) - Hunted to extinction by 1844 ? 9 Birds have big, complex brains! • Bird brains are larger relative to body size than all other vertebrates except mammals • Large motor-control areas (cerebellum), optical centers and song-control centers (cerebral hemisphere) What is “intelligence”? 19th century modern view Intelligence is the ability to “learn how to learn” – to solve problems of a given class correctly when first confronted, and then to solve problems of a similar nature with increasing speed when they are encountered again. Insight involves the ability to envision one's actions and their consequences. Corvids have shown a marked ability to solve problems. perch string meat Raven (Corvus corax) Raven 10 Black-capped Chickadees and Nuthatches can distinguish empty sunflower husks from full ones by weight alone. ... but chickens, doves, cats and rabbits don’t do as well in tests that involve reasoning or learning. Blue Jays have learned to avoid toxic Monarch butterflies by watching other Blue Jays in adjoining cages. Crows and parrots often perform as well as dogs in tests that involve reasoning or learning ... Blue Jays can locate food better than cats or monkeys. Great Tits learned how to open milk bottles ... 11 ... and other species learned the behaviour by imitation. House Sparrows have learned to open automatic doors to grocery stores and cafés by hovering in front of the sensor. Blue Tit Great Spotted Woodpecker European Starling Parrots can learn 100’s to 1000’s of different calls Class Aves (birds) II Respiratory system Circulatory system Digestion and excretion Metabolism Thermoregulation Senses Intelligence Communication Migration Dispersal Reproduction How to identify birds Communication Signal – behaviour that alters the behaviour of the receiver in a way that benefits the sender Display – ritualized signal that conveys a specific message Communication • Passive communication: plumage patterns – colors – shape – pattern • Active communication: – courtship displays – territorial displays – sound • calls • songs • non-vocal sounds 12 Passive communication: plumage patterns colors shape pattern Passive communication: plumage patterns colors shape pattern Grouse male vs female young vs adult Passive communication: plumage patterns colors shape pattern (incl. composition, number, size... Communication • Passive communication: plumage patterns – colors – shape – pattern • Active communication: – courtship displays – territorial displays – sound • calls • songs • non-vocal sounds Communication • Passive communication: plumage patterns – colors – shape – pattern Grouse displaying • Active communication: – courtship displays – territorial displays – sound • calls • songs • non-vocal sounds Mostly innate and highly stereotyped Communication • Passive communication: plumage patterns – colors – shape – pattern • Active communication: – courtship displays – territorial displays – sound • calls • songs • non-vocal sounds Red-winged Blackbird shoulder patch 13 Sound • Calls – short, stereotyped vocalizations; innate – Alarm – signals danger to other conspecifics – Contact calls – used to locate mates and young – Flight calls - keep the flock together Sound • Non-vocal sounds – – – – – Storks use bill clattering in courtship Mute swan hisses and grunts Woodpeckers peck to defend territories and attract mates Male grouse attract females by rapidly beating wings Broad-tailed hummingbirds trill with wings - territorial Migration Mass movement of individuals between breeding and wintering areas Defining characteristics: Sound Songs – complex, often learned – Territorial – Attract females – Species identification Class Aves (birds) II Respiratory system Circulatory system Digestion and excretion Metabolism Thermoregulation Senses Intelligence Communication Migration Dispersal Reproduction How to identify birds Not all long-distance movements are migrations e.g., Albatross fly huge distances when foraging - not seasonal or predictable - solitary • Seasonal (predictable) • Large numbers of individuals (flocks) 14 Not all long-distance movements are migrations Movement away from natal area: dispersal - solitary - new location unpredictable Not all long-distance movements are migrations Movement away from natal area: dispersal - solitary - new location unpredictable Movement away from a food-poor area: irruptions - not annual - unpredictable Migration - 75% of North American birds migrate. - 90% of “Canadian” birds migrate outside of Canadian borders - Migratory restlessness – controlled by hormones activated by changing daylengths - Many individuals use flyways, but these can change and some individuals do not “follow the pack”. Why do birds migrate? Birds migrate when benefits are higher than costs (energy and mortality risk) of using well-separated breeding and wintering areas. http://www.birdnature e.com/allflyways.html Snowy Owl Home | Site Introduction | Survival Needs | Bird Identifications Gallery & Profiles | Habitats of Birds | Bird Migration | Odds & Ends | Birds and Nature Magazine | Resources | Search Why do birds migrate? • Some species migrate south in the winter because food is unavailable in temperate zone • Higher reproductive success in temperate vs tropical areas Temperate zone: Abundant food and longer days for foraging in spring/summer Food shortages and cold in winter Tropics: High density of nest parasites and predators Mild temperatures and food availability in winter 15 Modes of Migration • • • • • Flight pattern Time of day Flight mode Length of migration Completeness Modes of Migration • Time of day Modes of Migration • Flight pattern – nonstop flights over oceans, deserts, mountains. – e.g., American Golden Plovers fly 4,000 km between NE U.S. and South America – in bad weather, birds can be grounded for hours or days (fallouts) – “hops” of 250-300 km Modes of Migration • Flight mode – diurnal migrants • often depend on thermal winds or updrafts, e.g., hawks. • may depend on diurnal food resources, e.g., swallows – nocturnal migrants • start about ½ hour after sunset, e.g., woodpeckers, vireos, thrushes • cooler and air less turbulent • avoid diurnal predators – soarers • flight concentrated in mid-day when thermals occur, e.g., hawks – powered flight • often crepuscular (start just before dawn) • nocturnal migrants Why fly at night? Modes of Migration • Length of migration – long-distance migrants • usually between continents • e.g., Golden Plover (between NE U.S. and S.A.) Modes of Migration • Completeness –complete migrants • entire population migrates –partial migrants – short-distance migrants • within continents • often elevational changes, e.g., Hermit Thrush • some individuals overwinter on the breeding grounds 16 Modes of Migration • Flight pattern How do they do it? • Orientation – ability to align in an appropriate direction when released into unfamiliar surroundings (internal compass). – nonstop – “hops” • Time of day – diurnal – nocturnal • Flight mode – soarers – powered flight • Navigation – ability to find a specific geographical location from a known starting point. • Length of migration – long distance – short distance • Completeness – complete migrants – partial migrants Orientation • Sun compass – can tell direction from the Sun’s position because of an internal circadian clock. Polarization pattern – e.g., at 7:00 AM a bird “knows” the Sun should be in the east, a few degrees above the horizon. – also, many birds can detect the polarization pattern, tt and d this thi can be b used d on days d with ith significant cloud cover. • Star compass – the north star (Polaris) gives an absolute point of reference. • Magnetic compass – can detect the Earth’s magnetic field. Orientation • Sun compass – can tell direction from the Sun’s position because of an internal circadian clock. – e.g., at 7:00 AM a bird “knows” the Sun should be in the east, a few degrees above the horizon. – also, many birds can detect the polarization pattern, tt and d thi this can be b used d on days d with ith significant cloud cover. • Star compass – the north star (Polaris) gives an absolute point of reference. • Magnetic compass – can detect the Earth’s magnetic field. 90° to Sun Navigation • Navigational landmarks – coastline – mountain ranges – large rivers • Magnetic anomalies • Local landmarks 17 Class Aves (birds) II Respiratory system Circulatory system Digestion and excretion Metabolism Thermoregulation Senses Intelligence Communication Migration Dispersal Reproduction How to identify birds Dispersal Natal dispersal – the first movement away from the nest area (typically, high mortality). B Breeding di dispersal di l – movements t off adults between different breeding locations. Amniotic egg Dispersal Not all birds migrate, but almost all individuals disperse. There are two types of dispersal …. Class Aves (birds) II Respiratory system Circulatory system Digestion and excretion Metabolism Thermoregulation Senses Intelligence Communication Migration Dispersal Reproduction How to identify birds Amniotic egg 18 Reproductive System Males • testes small, bean shaped most of year • testes enlarge 300 x at breeding season • sperm stored in seminal sac • most males lack a penis Birds have a single ovary Females • only left ovary and oviduct develop • eggs fertilized at top of oviduct • albumin, shell membrane, shell added during passage down oviduct Reproductive behaviour Monogamy Most birds are monogamous for one or more breeding seasons and some are monogamous for life (e.g., swans and geese) • Monogamy • Individually defended territories -most birds parakeets mating • Colonial breeding - about 13% of all bird species • Cooperative C ti b breeding di – about b t 250 species i • Non-monogamous breeding (<10% of bird species) Two kinds of monogamy: Social monogamy – male and female cooperate in raising young; >90% of all birds. Genetic monogamy – male and female are the sole genetic parents of all the young; rare. rare Birds can be either socially or genetically monogamous, both – or neither. • Brood parasitism • Courtship ritual - both sexes participate Social vs genetic monogamy implies … Extra-pair copulations Why? • forced copulations – waterfowl • females are often willing and actively seek extra-pair copulations - hedge against infertility in primary partner - increases genetic diversity of young - might produce young by a higher-quality male How lack of genetic monogamy is detected If the DNA of young and putative father do not match extra-pair copulation If the mtDNA of young and mother do not match The bottom line is …. Genetic monogamy is probably rare nest parasitism (egg dumping) 19 Reproductive behaviour • Monogamy • Individually defended territories - most birds • Colonial breeding - about 13% of all bird species • Cooperative C ti b breeding di – about b t 250 species i • Non-monogamous breeding (<10% of bird species) Individually defended territories • most territories contain both food and nesting sites • some territories contain only nesting sites • boundaries are dynamic • males l typically t i ll defend d f d territories t it i by b – song – visual display behaviour – active defense • Brood parasitism • Courtship ritual - both sexes participate Reproductive behaviour • Monogamy • Individually defended territories - most birds • Colonial breeding - about 13% of all bird sp. What are the advantages and disadvantages of colonial breeding? Reproductive behaviour • Monogamy • Individually defended territories - most birds • Colonial breeding - about 13% of all bird sp. Colonial Breeding Advantages: - favored by scarcity of sites safe from predators - favored by scarcity of sites near abundant food - cooperative protection from predators - enhances foraging on ephemeral food supplies Disadvantages: - colony may attract predators - increase the spread of nest parasites (e.g., mites, lice) - increased competition for food Cooperative breeding • breeders defend group territories • nest helpers – young birds that help feed and defend the nest • nest helpers benefit by - raising close relatives (kinship selection) - creating large territories in which they might breed • Cooperative breeding – about 250 sp. (< 3%) Florida Scrub Jay 20 Reproductive behaviour • Monogamy Non-monogamous breeding • Promiscuity – no stable social relationship • Individually defended territories -most birds – rare in birds; found in some hummingbirds • Colonial breeding - about 13% of all bird sp. • Cooperative breeding – about 250 sp. (< 3%) • Non-monogamous breeding (<10% of bird sp.) • Brood parasitism • Courtship ritual - both sexes participate Non-monogamous breeding • Promiscuity – no stable social relationship – rare in birds; found in some hummingbirds • Polygyny – one male and several females • Polyandry – one female and several males • Polygynandry – both males and females have multiple mates Polygyny (one male and several females) • 2% of all bird species • males provide little or no parental care • young are typically precocial – rare; Smith’s Longspur in North America Polygyny (one male and several females) • • • • 2% of all bird species males provide little or no parental care young are typically precocial lekking – males gather at the lek and display – females f l choose h males; l high-status hi h males l have h multiple mating – no male parental care Hatch blind, naked, and virtually helpless; most songbirds, hawks, herons, etc. Hatch with eyes open, downy feathers, and are capable of independent movement within a few hours; e.g., grouse, quail, ducks, water birds, shorebirds, etc. e.g. sage grouse 21 Non-monogamous breeding • Promiscuity – no stable social relationship – rare in birds; found in some hummingbirds • Polygyny – one male and several females • Polyandry – one female and several males • Polygynandry – both males and females have multiple mates – rare; Smith’s Longspur in North America Polyandry • Polyandry – one female and several males – 1% of all bird species – classic polyandry: female lays eggs in separate nests and males incubate and rear brood – cooperative polyandry: female lays eggs in one nest and several males care for young – reversed sexual dimorphism: females larger and more colorful than males Spotted Sandpiper Spotted Sandpiper (male) Non-monogamous breeding • Promiscuity – no stable social relationship – rare in birds; found in some hummingbirds • Polygyny – one male and several females • Polyandry – one female and several males • Polygynandry – both males and females have multiple mates – rare; Smith’s Longspur in North America Reproductive behaviour • Monogamy • Individually defended territories -most birds • Colonial breeding - about 13% of all bird sp. • Cooperative breeding – about 250 sp. (< 3%) • Non-monogamous breeding (<10% of bird sp.) • Brood parasitism Brood Parasitism • Females lay their eggs in the nests of other birds (the hosts) • Two types of brood parasitism: – intraspecific parasitism (“egg dumping”) • very common – interspecific parasitism • European cuckoos • North American Cowbirds • Courtship ritual - both sexes participate 22 Reproductive behaviour • Monogamy • Individually defended territories -most birds • Colonial breeding - about 13% of all bird species • Cooperative C ti b breeding di – about b t 250 species i • Non-monogamous breeding (<10% of bird species) • Brood parasitism • Courtship ritual - both sexes participate Blue-footed boobies and frigate birds Galapagos Islands 2006 Western grebe - weed dancing Western Grebe - rushing Nest sites trees e.g., Bald Eagle shrubs, e.g., Bushtit 23 Nest sites cliff ledge: e.g., Horned puffin Nest sites crevices or cliff faces - e.g., Cliff swallow on ground e.g., gulls, terns, loons, ducks Class Aves (birds) II Respiratory system Circulatory system Digestion and excretion Metabolism Thermoregulation Senses Intelligence Communication Migration Dispersal Reproduction How to identify birds treeholes e.g., woodpeckers Eight visual categories 1) 2) 3) 4) 5) 6) 7) Swimmers – ducks, etc. Aerialists – gulls, etc. Long-legged waders – herons, cranes, etc. Smaller waders – Plovers, sandpipers, etc. Fowl like birds – Grouse, Fowl-like Grouse quail quail, etc etc. Birds of prey – Hawks, eagles, owls Nonpasserine land birds – Kingfishers, woodpeckers, hummingbirds, swifts, etc. 8) Passerine (perching) birds What is the bird’s size? What shape is its tail? Use some common birds as a “yardstick”. rounded forked and pointed Robin Pigeon House Sparrow Quail Barn Swallow 24 What shape is its bill? What shape are its wings? Hawk Tern Sparrow Tern Hawk Warbler How does it behave? House Wren Does it climb trees? Flycatcher jerks down headfirst Woodpecker Nuthatch How does it fly? spirals Brown Creeper Does it swim? soar Up and down hover Mallard Loon Kingfisher Northern Flicker Hawk 25 Does it dive? Does it wade? Sandpiper Mallard Merganser Merganser Bird Topography - What are its field marks? Great Blue Heron What are its field marks? Color patterns i.e. tibia! Western Meadowlark What are its field marks? Color patterns Wood Duck What are its field marks? Color patterns American Wigeon 26 What are its field marks? Tail patterns Western Kingbird Eastern Kingbird What are its field marks? Tail patterns What are its field marks? Tail patterns Western Kingbird What are its field marks? Tail patterns Rufous-sided Towhee Dark-eyed Junco What are its field marks? Tail patterns American Redstart What are its field marks? Rump patches Cliff Swallow 27 What are its field marks? Eyestripes and eye-rings Mountain Chickadee Black-capped Chickadee What are its field marks? Wing bars Ruby-crowned kinglet What are its field marks? Wing Patterns Herring gull 28